Revision of zamia in mega mexico

8/10/2019 Revision of zamia in mega mexico

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Taxonomic revision ofZamiain Mega-Mexico

FERNANDO NICOLALDE-MOREJN1, ANDREWP. VOVIDES1,

ANDDENNISW. STEVENSON2

1 Departamento de Biologa Evolutiva, Instituto de Ecologa, A. C. km 2.5 Antigua Carretera aCoatepec No. 351, Xalapa 91070 Veracruz, Mexico

2 The New York Botanical Garden, Bronx, NY 10458-5120, USA; e-mail: [email protected]

Abstract. The genusZamiais revised for Mega-Mexico, with 22 species recognizedand described. The study presents a taxonomic clarication for the genus in Mesoa-merica, a contribution that provides the foundation for a future monograph forZamiain the Neotropics. The largest proportion of species richness and endemism for the

genus is concentrated in southeastern Mexico, among the states of Chiapas, Oaxaca,Tabasco, and Veracruz, an area that is considered highly diverse in oristic terms.Distribution maps and a key to species are also provided, as well as complete des-criptions of the specimens examined, including information on nomenclatural types,habitats, synonymies, and etymologies. A lectotype is designated forZamia loddigesii,and neotypes forZ. galeottii,Z. leiboldiivar.angustifolia, andZ. variegata.Zamia

sparteais illustrated for the rst time, and chromosome numbers forZ. herrerae arereported and illustrated. Finally, scanning electron micrographs of leaet trichomecharacter states are presented, along with a discussion of their systematic implicationswithin the group.

Key Words:Endemism, Mexico, gymnosperms, cycads, oristic richness,Zamia.

Resumen. El gneroZamiaes revisado para Mega-Mxico, con 22 especies descritasy una species dubia. El estudio est orientado al esclarecimiento taxonmico delgnero en Mega-Mxico, una contribucin que siente los fundamentos para una futuramonografa para Zamia en el Neotrpico. La mayor riqueza y endemismo para elgnero se concentra en el sureste de Mxico, entre los estados de Chiapas, Oaxaca,Tabasco y Veracruz, rea de alta biodiversidad orstica. Mapas de distribucin y unaclave para las especies son presentadas, como tambin descripciones completas, tipos,hbitat, sinnimos, etimologa y especimenes examinadas. Se designa un lectotipo

paraZamia loddigesii, y neotipos paraZ. galeottii,Z. leiboldiivar.angustifoliayZ.variegata.Zamia sparteaes ilustrada por primera vez y nmeros cromosmicos paraZ. herreraeson presentados. Finalmente, se presentan fotografas de tricomas al mi-croscopio electrnico y una discusin de sus implicaciones en la sistemtica delgrupo.

According to Stevenson (1992), Zamiaceaecomprises eight genera distributed in tropicaland subtropical Africa, Australia, GreaterAntilles, North, Central and South America.Five genera with 94 species are known fromthe Neotropics (Hill et al., 2007). The genera

Ceratozamia Brongn. (21 spp.) and DioonLindl. (13 spp.) are both endemic to Mexicoand a neighboring biogeographic region of

Central America that is oristically similar tothe southern part of Mexico (Mega-Mexico)whileMicrocycasA. DC. (1 sp.) andChiguaD.W. Stev. (2 spp.) are endemic to Cuba andColombia, respectively. The type genus forthe family, Zamia L. (59 spp.), is distributed

throughout tropical and sub-tropical Americaas well as the Caribbean, with exception tothe Lesser Antilles.

Brittonia, 61(4), 2009, pp. 301335 ISSUED: 1 December 2009 2009, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.


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Zamiais the widest distributed genus of theorder Cycadales in the Neotropics. Its northernrange starts in Georgia and Florida (U.S.A.),reaching Bolivia and the Mato Grosso ofBrazil in South America (Balduzzi et al.,1982; Sabato, 1990; Norstog & Nicholls,1997; Stevenson, 2001a, b). A remarkablemorphological and cytological variation has

been documented (Vovides,1983; Moretti &Sabato,1984; Moretti,1990a,b; Stevenson etal., 19951996a; Vovides & Olivares, 1996;

Norstog & Nicholls, 1997), and also highlevels of genetic variation (Gonzlez-Astorgaet al., 2006). As a consequence of thiscomplexity, the taxonomy ofZamiais contro-versial; although the genus comprises 75

species, their circumscription and limits haveyet to be determined (Hill et al.,2007).The most recent exhaustive taxonomic

treatment forZamiawas published by Schus-ter (1932) and later work by Sabato (1990)and Stevenson(1987;1991a,b;1993;2001a,

b;2004) underlined nomenclatural and taxo-nomic anomalies inZamia, principally owingto insufcient eldwork and a scarcity ofgood-quality botanical collections. The taxo-nomic history for the genus in Mexico began

with the publication ofZamia fufuraceaL. f.in 1789 from the central-south coastal regionof Veracruz, which also represents the rstcycad species described from the Americancontinent. During the 19th century, six spe-cies were subsequently described for Mexico;Z. scheriMiq.,Z. katzeriana(Regel) Rettig,Z. lawsoniana Dyer, Z. loddigesii Miq., Z.

spartea A. DC. and Z. verschaffeltii Miq.,whereas the remaining known species weredocumented and characterized during the20th century, with a marked tendency fortaxonomic activity during the last thirty years(Vovides et al.,1983; Schutzman et al.,1988;Stevenson et al., 19951996a, b; Schutzman& Vovides,1998; Vovides,1999).

The present taxonomic revision includesendemic species of the cycad genusZamiathatoccur in Mega-Mexico 2, a term coined byRzedowski (1991) that associates the CentralAmerican territories of Guatemala, Belize,Honduras and northern Nicaragua to theMexican states of Nayarit on the Pacic to

southern Tamaulipas on the Gulf of Mexico usingbiotic (mainly oristic) criteria. Rzedowskisconcept will be referred to as simply Mega-

Mexico hereafter, given that the slight dif-ferences in the boundaries of Mega-Mexico 1with respect to Mega-Mexico 2 do not affectthe biogeographic aspects of our descriptionof the species. This study is intended to

provide a taxonomic clarication of the genusin Mesoamerica and to provide the basis for afuture monograph ofZamia.

Materials and methods

The present taxonomic revision is based onmore than 450 specimens from the followingherbaria: B, BM, CIB, CHIP, CICY, ECOSUR,ENCB, F, FCME, FLAS, FTG, HEM, IBUG,

IEB, K, LE, MEXU, MO, NY, SERO, U,UADY, UAMIZ, US, W, WIS, XAL, XALU,and ZEA. Unfortunately, we were not able toobtain vouchers on loan from NAP; therefore,material from this herbarium is not cited.

Chromosome counts for Zamia herreraeCaldern & Standley were performed on veindividuals held at the Jardn Botnico Fco.Javier Clavijero, Instituto de Ecologa, A.C.(JBC). These plants has been previouslycollected at the Acacoyagua and Tonal

regions, located in the state of Chiapas andrepresent the species range in southern Mex-ico. Plants from its full biogeographic range,which would include El Salvador and Guate-mala, were not available for this study. Amodied root-tip squash method was used forexamining somatic metaphase cells described

by Vovides (1983) with a 12 to 15 hour ice-water (0C) follow-up soak after the 0.2%colchicine pretreatment at ambient tempera-ture (Schutzman et al., 1988). Counts weremade from the best 1015 metaphase cellsand karyotype noted according to the classi-cation of Schlarbaum and Tsuchiya (1984).Photomicrographs were produced using aZeiss Fomi III photomicroscope tted with

planapochromatic objectives and Kodak Plus-X pan ASA 125 lm.

Scanning electron micrographs (SEM)were taken on young leaet material fromliving plants cultivated in the JBC. Sampleswere placed on sample stubs with doublesided adhesive tape and then introduced into

a dessicator for 24 hours. All samples weresputter coated with gold-palladium at 1.5 kvat 5 mA for 8 minutes with a Jeol Fine Coat

302 BRITTONIA [VOL 61


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JFC 1100 sputter coater. Observations weremade with a Jeol JSM-5600LV SEM.

In all cases, the types have been examinedby one or more of the authors.

ResultsHABITAT

Of the 22 species included in this revision,18 occur in specic habitats, accounting forthe restricted distribution of the majority ofthe taxa. The species with the widest distri-

bution are associated with two or morevegetation types, namely (a)Zamia paucijugaWieland, found in pine-oak, oak and tropicaldry forests; (b)Z. polymorphaD.W. Stev., A.

Moretti & Vzq. Torres, located in evergreentropical rainforest, sub-deciduous tropicalforest and their secondary succession stages;(c) Z. herrerae, which occurs in evergreentropical rainforest, sub-deciduous tropicalforest and tropical dry forest and theirsecondary succession stages and nally, to alesser extent; and (d)Z. loddigesii, present inevergreen tropical rainforest, but more com-monly in sub-deciduous tropical forest and itssecondary succession stages.

MORPHOLOGY

HABIT.All adult cycad stems are pachycau-lous and may be columnar and arborescent orsubterranean and tuber-like. The generaDioon,

Microcycas, Ceratozamia and LepidozamiaRegel are usually columnar arborescent in habit,while the subterranean forms are characteristicofBowenia Hook. ex Hook. f., Chigua, andStangeria T. Moore. Cycas L., EncephalartosLehm.,MacrozamiaMiq. andZamiahave bothstem morphologies, either subterranean tuber-like or columnar arborescent (Stevenson,1980).In this context, the species of Zamia inMesoamerica represent both growth forms, withthe subterranean tuber-like habitat predominant.Only four species,Z. inermis,Z. onanreyesii,Z.

soconuscensis andZ. tuerckheimii, have arbo-rescent stems reaching up to 100 cm or more inheight. Some species branch dichotomouslywith age (namely, Z.scheri Miq., Z. furfuracea,Z. inermis Vovides, Rees & Vzq. Torres, Z.

soconuscensis Schutzman, Vovides & Dehgan,Z. loddigesiiandZ. herrerae), with the coastaldune species Z. furfuracea being the most

notable, with branches reaching up to 80 cmlong in adult plants.

TRICHOMES.Trichomes of cycads leavesare bi-celled, consisting of a small basal celland a longer free apical portion (Stevenson,

1981). All the trichomes analyzed here showthe same bifurcate pattern, with one arm

proportionally longer than the other (Fig.1A).Each trichome presents a rounded basal celland a more extensive cylindrical bifurcate free

portion, which in most cases observed hadevidence of collapse and twisting (Fig.1BF).The pubescence was signicantly denser inemerging than in older adult leaves with thelatter becoming completely glabrous. Anexception to this condition is found in Zamia

furfuracea, which maintains a great part of itsoriginal indument on the abaxial surface ofeach leaet.

According to Stevenson (1981), four typesof trichomes occur in Zamia; of these, thetransparent ramied and the colored ramiedare the types found in the present work. Incontrast to what was found in the aforemen-tioned study by Stevenson, no trend orcorrelation was found between aerial stems(i.e., Z. soconuscensis or Z. inermis) and

bifu rcat e tr ic home s wi th equa l le ngthbranches. Trichomes with unequal sizedprevail among taxa with both aerial andsubterranean stems. In this context, weconsider that a more extensive and detailedsampling of the genus Zamia across the

Neotropics would be necessary to corroborateany correlation between trichome morpholo-gy and stem habit.

REPRODUCTIVE STRUCTURES.Although thecharacters employed for the identication ofspecies of Zamia have been obtained fromleaf morphology (Miquel,1861,1869; Regel,1857, 1876; de Candolle, 1868; Schuster,1932; Eckenwalder, 1980; Vovides et al.,1983; Newell, 1986; Schutzman & Vovides,1998; Schutzman et al., 1988; Stevenson,1993, 2001a, b, 2004; Nicolalde-Morejn etal., 2008), the evaluation of reproductivecharacters, especially those corresponding toovulate strobili, is essential to discriminateamong closely related taxa. Outstandingattributes that should be considered in this

case are (i) the form and shape of the coneapex (Stevenson, 1987); (ii) the peduncle

position with respect to the vertical axis of

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the cone, when mature (Schutzman et al.,1988); and (iii) the overall color of the cone(see descriptions and Figs.2,3). In contrast tothe ovulate strobili, pollen strobili showscarce variation at the species level, and theirutility to discriminate among species that

might possess high degrees of genealogicalafnity is relatively low. For the descriptionof the pollen reproductive axes, the terminol-

ogy introduced by Mundry and Sttzel (2003)has been followed.

CHROMOSOME NUMBERS.Zamia shows thehighest chromosome numbers and karyotypevariation throughout the order Cycadales,with 2n counts of 16, 17, 18, 22, 23, 24, 25,

26, 27, and 28 (Marchant, 1968; Norstog,1980, 1981; Vovides, 1983; Moretti &Sabato, 1984; Moretti, 1990a, b; Vovides &

FIG. 1. Trichomes.AB. Zamia furfuracea.CD.Z. katzeriana.EF.Z. polymorpha.



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FIG. 2. Illustrations of ovulate strobili.A.Zamia inermis.B.Z. vazquezii. C. Z. paucijuga.D.Z. lacandona.E.Z.spartea.



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FIG. 3. Illustrations of ovulate strobili.A. Z. scheri.B. Z. furfuracea.C. Z. cremnophila.D. Z. katzeriana.E. Z.polymorpha.



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Olivares,1996). In contrast, all Ceratozamiaspecies studied so far have stable diploidchromosome numbers (2n=16) and karyo-types, as do all the Dioonspecies analyzed todate with stable diploid chromosome numb-

ers (2n=18) and karyotypes (Marchant,1968;Vovides,1983,1985; Moretti,1990a,b).

Chromosome counts are presented andillustrated (Fig. 4) here for the rst time forZamia herrerae. This species has, but to alesser extent, cytotype polymorphisms similarto those found for Z. paucijuga (Moretti &Sabato,1984) andZ. polymorpha(Vovides &Olivares, 1996; Stevenson et al., 19951996b).Zamia herrerae has 2n=23, 24 fromtwo populations along its Mexican range in

Chiapas (Fig.4). Both the m (median regionof the chromosomes) and T (telocentric)chromosomes vary in number (611 T, 46 m) and are large (611m for T and 812m for m) and their arms can be longer thanhalf the spindle axis, which can cause mitoticinstability during telophase (Schubert, 2007).Karyotype differences are probably due tocentric ssions occurring on some of thelarger m chromosomes, giving rise to telo-centrics with part of the centromere still

present. In this context, there are two generalhypotheses to explain karyotypic evolution in

Zamia: rst, Norstogs hypothesis (Norstog,1980,1981) relating karyotype simplicationand symmetry with progressive fusion oftelocentric chromosomes, which predicts lowdiploid number in taxa with a high number of

metacentric chromosomes, and secondly thatof more recent research on Zamia, which

postulates centric ssion rather than fusionproducing a progressively higher diploid num-ber and asymmetric karyotypes with a highnumber of telocentric chromosomes (Moretti& Sabato, 1984; Vovides & Olivares, 1996;Caputo et al., 2004). For a more in-depthdiscussion of mechanisms of chromosomeevolution in seed plants, see Jones (1998).

The highly asymmetric karyotypes and

somatic chromosome numbers in both Z.paucijuga and Z. polymorpha, which alsoappear to be the pattern forZ. herrerae, seemto be correlated with the highest morpholog-ical variation and widest geographic distribu-tion of the genus in Mesoamerica. Zamiaherrerae has a range of about 1000 km,spanning El Salvador, Guatemala, and Chia-

pas (Mexico); therefore, we suggest investi-gating the distribution of chromosomecharacter states in this species throughout its

range. This karyotype asymmetry contrastswith their congeners of a more restricted

FIG. 4. Chromosomes ofZamia herrerae.A. 2n= 23.B. 2n= 24.



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distribution, which often have less morpho-logical variation and a tendency towardsconstant chromosome number and karyotype,e.g., Z. cremnophila Vovides, Schutzman &Dehgan,Z. scheri,Z. inermis,Z. katzeriana,

Z. purpureaandZ. soconuscensis(all 2n=16),as well as Z. furfuracea, Z. spartea and Z.vazquezii (all 2n=18). Vovides and Olivares(1996) and Jones (1998) comment thatatypical chromosome number increase attrib-uted to ssion is probably a result of stressfulinuences.

DISTRIBUTION ANDENDEMISM

Seventy-ve percent of the species in this

revision are endemic to the type locality andnearby areas. They are limited to two or threeclose populations with low population densi-ties. These attributes are consistent withRabinowitzs (1981) evaluation criteria forspecies rarity, which mainly considers infor-mation related to geographic range, habitatspecicity and local population size. Incongruence with these criteria, the endemicZamiaspecies of Mesoamerica are consideredrare, threatened or endangered and are listed

under the IUCN Red List (IUCN,2005; Hillet al.,2007).With 21 endemic cycad species in three

genera, Mexico has the highest cycad diver-sity and number of endemics of the region.Six species of Zamia are known in Guate-mala, two of which are endemic (Z. monticolaChamberlain and Z. tuerckheimii J. DonnellSmith); in Honduras, three species are knownof which three recently described species areendemic (Z. oreillyi C. Nelson, Z. sandovaliiC. Nelson, andZ. onanreyesiiC. Nelson & G.Sandoval). Two further species are known forBelize, of which one is endemic (Z. prasinaW. Bull) and nally, El Salvador is repre-sented by one broadly distributed species, Z.herrerae, with a range that runs along thePacic seaboard through Guatemala and theSierra Madre de Chiapas in Mexico. Otherspecies of the genus in Mexico with broaddistributions are (a) Z. paucijuga, distributedalong the Pacic seaboard of Mexico rangingfrom Nayarit (northwest Mexico) to Oaxaca

(southwest); (b) Z. polymorpha, distributedwidely throughout the Yucatn penninsula inMexico, Belize, and the Petn region of

Guatemala; (c) Z. loddigesii, ranging alongthe Gulf of Mexico seaboard from Tamauli-

pas (northeast Mexico) to Tabasco (south-east); and (d)Z. variegataWarsz., distributed

between Guatemala and southern Mexico.

The southern and southeastern regions ofMexico, comprising the states of Veracruz,Oaxaca, Chiapas, and Tabasco, are the mostdiverse area of Mexico for the genus Zamia,with seven endemic species (Z. cremnophila,Z. lacandona,Z. loddigesii,Z. katzeriana,Z.

purpurea, Z. polymorpha, and Z. spartea).The Gulf of Mexico region has three micro-endemics Z. furfuracea, Z. inermis and Z.vazquezii, whereas two micro-endemic spe-cies, Z. scheri and Z. soconuscensis, are

known respectively from the Sierra MadreOriental and the Sierra Madre de Chiapas.Finally, in spite of the status of Z. pauci-

jugaas a Mexican endemic, its distribution isextremely wide within Mega-Mexico, cover-ing a range of about 1000 km between thestates of Oaxaca and Nayarit. The northernlimit of Z. paucijuga in the latter staterepresents the northernmost distribution forthe genus along the Pacic seaboard of the

Neotropics.

Taxonomic treatment

Zamia L., Sp. Pl. ed. 2: 1659. 1763. nom.cons. Type:Zamia pumilaL.

Palma-FelixAdanson, Fam. Pl. 2: 21, 587.1763.

AulacophyllumRegel, Gartenora 25: 140.1876.

Stems hypogeous and epigeous, erect todecumbent, sometimes dichotomously

branched in mature plants.Cataphyllscharta-ceous to membranaceous, stipulate, persistentor deciduous, base triangular, apex longacuminate to aristate, tomentose, generallyreddish-brown to yellowish. Ptyxis erect toinexed. Leaves stipulate, ascending todescending to spreading, reddish-brown orgreen when emerging; petiole sometimes

blackish in young leaves, terete or subterete,without prickles or heavily to lightly armedwith straight or bifurcate prickles; rachis

subterete generally with few prickles alongthe proximal third or without prickles, withup to 60 pairs of leaets. Leaets articulate,



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sessile, papyraceous to coriaceous, linear,linear-lanceolate, lanceolate, ovate, obovateobpyriform to elliptic, opposite to suboppo-site, falcate or non-falcate, imbricate tonon-imbricate, generally acute at apex and

symmetric, attenuate at base, margins entireto dentate along upper third, subrevolute,articulations green, yellowish or dark brownin young leaets. Pollen strobili usually 12(4), with sterile tip, erect, cylindrical toconical, light brown to purple, tomentulose,apex acute to apiculate, generally with dense-ly tomentose peduncles; pollen sporangio-

phores cuneiform, distal face truncate

hexagonal, 0.30.55 cm long, fertile abaxialsurface 2 lobed with 214 bisporangiatesynangia per lobe, sporangia dehiscent bylongitudinal slit. Ovulate strobili usuallysolitary, erect to decumbent, cylindrical to

ellipsoid, purple to yellowish, generallytomentulose, apex acute to apiculate; pedun-cle densely tomentose; ovulate sporangio-

phores cuneiform-peltate to scutiform, distalend truncate-hexagonal when not scutiform.Seeds ovoid, sarcotesta white to pink whenimmature, red at maturity, sclerotesta smooth

but sometimes with several furrows runninglongitudinally from micropylar end.

Key to the species ofZamiain Mega-Mexico1. Leaets chartaceous to papyraceous.

2. Leaet margin dentate.3. Leaets elliptic, adaxial surface with yellow to cream variegation . . . . . . . . . . . . . . . . . . . . . Z. variegata3. Leaets long-lanceolate, without variegation.

4. Leaets imbricate, peduncle of pollen strobili decumbent, up to 16 cm long. . . . . . . . . . . . . . Z. oreillyi4. Leaets not imbricate, peduncle of pollen strobili erect, up to 8 cm long . . . . . . . . . . . . . . . Z. herrerae

2. Leaet margin serrulate to entire.5. Leaet margin entire, chartaceous.

6. Leaets oblong-lanceolate, glossy, 46 cm wide; ovulate strobili iridescent blue-green at matu-rity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. tuerckheimii

6. Leaets linear-lanceolate, not glossy, 0.61.5 cm wide, ovulate strobili dark-brown at matu-rity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. soconuscensis

5. Leaet margin serrulate, papyraceous.7. Leaets sub-falcate basally, long-acuminate, strongly apically curved. . . . . . . . . . . . . Z. monticola7. Leaets straight, acuminate, not curved at apex.

8. Leaets elliptic to lanceolate, ovulate strobili cylindrical to ovoid, dark-green and glabrous whenmature; from San Lus Potos and Quertaro, Mexico . . . . . . . . . . . . . . . . . . . . . . . Z. scheri

8. Leaets ovate to obpyriform, ovulate strobili ovoid-cylindrical, gray to brown tomentulose whenmature; endemic to Veracruz, Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. vazquezii

1. Leaets coriaceous.9. Leaet margin entire to serrulate.

10. Leaet margin entire, petiole unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. inermis10. Leaet margin serrulate, petiole prickly.

11. Leaets long acuminate apically; stems arborescent . . . . . . . . . . . . . . . Z. onanreyesii11. Leaets rounded to acute apically; stems subterranean.

12. Leaets obovate to oblanceolate, keeled adaxially, apex rounded. . . . Z. furfuracea12. Leaets linear to oblanceolate, at adaxially, apex acute.

13. Leaets linear, 0.40.6 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . .Z. spartea13. Leaets lanceolate, 0.7 cm wide.

14. Leaets falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. sandovallii14. Leaets not falcate.

15. Leaets lanceolate-oblanceolate, ovulate strobili dark-brown tomentu-lose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. polymorpha

15. Leaets linear-lanceolate, ovulate strobili brown to yellowish.16. Ovulate strobili ellipsoid to cylindrical, apex acute to apiculate,

yellowish-brown; from the Pacic seaboard of Mexico . . . Z. paucijuga16. Ovulate strobili ellipsoid to conical, apex acute, yellowish; from the Gulf

of Mexico seaboard . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. loddigesii9. Leaet margin distinctly dentate.

17. Leaets channeled adaxially between veins, appearing plicate . . . . Z. purpurea17. Leaets smooth, not channeled adaxially between veins, not appearing plicate.

18. Leaets linear-lanceolate, imbricate, petiole strongly armed with straight orbifurcate prickles up to 6 mm long . . . . . . . . . . . . . . . . . .Z. cremnophila



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18. Leaets lanceolate to oblanceolate, not imbricate, petiolearmed with smallstraight prickles, generally between 24 mm.19. Leaets with brilliantly shining cuticle on adaxial surface;ovulate strobili

decumbent when mature. . . . . . . . . . . . . . . . . . . . . . . . Z. katzeriana19. Leaets without brilliantly shining cuticle on adaxial surface; ovulate

strobili erect when mature.

20. Distal leaets sub-falcate, petiole with bulbous base, blackish in youngleaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. lacandona20. All leaets straight, petiole with no bulbous base, greenish in young

leaves.21. Leaets oblong to lanceolate, bright grass-green, without conspic-

uously denticulate along margins; ovulate strobili green glabrouswhen mature, apex acuminate . . . . . . . . . . . . . . . . Z. prasina

21. Leaets long-lanceolate, conspicuous dentate, up to 4 mm long;ovulate strobili brown tomentulose when mature, apex long-apiculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. standleyi

Zamia cremnophila Vovides, Schutzman &

Dehgan., Bot. Gaz. 149(3): 351. 1988. Type:Mexico. Tabasco: 18 Aug 1981, M. A.Magaa&S. Zamudio 343 (holotype: MEXU;isotypes: CSAT, FCME, MO, UAMIZ).

Stem hypogeous, generally unbranched, 534 cm long, 411.3 cm in diam. Cataphyllschartaceous, persistent, base triangular, apexaristate, 5.4 cm long, 2.9 cm at base, reddish-

brown tomentose. Ptyxis inexed. Leaves 23(4), 57.517940.276.1 cm, descending, red-dish-brown when emerging; petiole 10.295.3 cm long, blackish in young leaves,subterete, heavily armed with straight or bifur-cate prickles up to 6 mm long; rachis subterete,up to 84 cm long, with few prickles along the

proximal third. Leaets 528 pairs, sessile,coriaceous, lanceolate, opposite to subopposite,imbricate, apex acute, base attenuate, marginsdentate along distal third, subrevolute; articula-tions dark brown in young leaets, 0.81.2 cm;the median leaets 22.7383.14.4 cm wide.

Pollen strobiliusually 12, erect, cylindrical, upto 8 cm long and 1.4 cm in diam, light brown,apex acute; peduncle densely light browntomentose, 4.5 cm long, 1.2 cm in diam; pollensporangiophores cuneiform, distal face truncate-hexagonal, 0.35 cm long, fertile abaxial surfacewith 3 bisporangiate synangia per lobe.Ovulate

strobiliusually solitary, erect, ellipsoid, 13.2 cmlong, 4.9 cm in diam, brown to reddish,tomentulose, apex acuminate; peduncle densely

brown tomentose, 2.9 cm long, 1.6 cm in diam;

distal face hexagonal-truncate with a horizontallongitudinal depression, 11.2 cm high, 1.11.4 cm wide. Seeds ovoid, sarcotesta white

when immature, red at maturity, 22.8 cm long,

1.4

2 cm in diam, sclerotesta smooth.Chromosome number.2n=16 (Schutzmanet al.,1988).

Diversity and genetic structure.The av-erage of alleles per locus is A=1.98, the

percentage of polymorphic loci is P= 94.3, theexpected heterozygosity is HE=0.347 andthe genetic differentiation between the two

populations currently under study is Fst=0.093 (Gonzlez-Astorga et al. unpubl. data).

Distribution and habitat.Endemic toTabasco, Mexico (Fig.5), on karstic rocks andcliffs of the Sierra El Madrigal, between 50150 m. The vegetation type where this speciesgrows is evergreen tropical rain forest orbosquetropical perennifolioof Rzedowski (1978).

Etymology.The specic epithet is derivedfrom the Greek word for cli ff-friend( = cremnos = cliff, and =los = friend/lover; Schutzman et al., 1988),

because of its unusual habitat.Distinguishing features.The species is

characterized by its exclusive habit on the

rocky walls of limestone cliffs, in addition todescendent leaves. Petioles are densely armedwith prickles that are sometimes branched,and also lanceolate, imbricate leaets that arevisibly dentate along the distal third.

Additional specimens examined. MEXICO.TABASCO: Teapa, F. Nicolalde-Morejn et al. 1497(XAL), M. A. Prez-Farrera 293 (HEM), M. A. Prez-

Farrera 900 (HEM, MEXU).

Zamia scheriMiq. in Lem., Hort. Vanhoutt. 1:

20. 1845. Type: ex Horto Petropolitano in H.Houtte. vecta,Miquel s.n. (neotype, designatedby Stevenson & Sabato,1986: U). (Fig.7)



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Stem subterranean, dichotomously branchingin older plants, up to 30 cm long, 48 cm indiam. Cataphylls membranaceous, persistent,

base triangular, apex aristate, 4.5 cm long,1.2 cm at base, yellowish tomentose. Ptyxisinexed. Leaves 15(8), 1545820 cm,ascending to spreading, dark-brown whenemerging; petiole 914 cm long, blackish inyoung leaves, unarmed, terete; rachis subterete,up to 31 cm long, unarmed. Leaets 2035

pairs, sessile, papyraceous, elliptic to oblanceo-late, alternate to subopposite, apex acute sym-metric, base cuneate, margins serrulate alongdistal third, subrevolute; articulations light-

brown in young leaets, 0.30.4 cm wide,median leaets 591.54.5 cm. Pollen stro-bili usually 13, erect, conical, 46 cm long,1.52.2 cm in diam, gray tomentose, apexacute; peduncle yellowish tomentose, 2.8 cmlong, 0.91.1 cm in diam; pollen sporangio-

phores cuneiform, distal face hexagonal-

truncate, 0.3 cm long, fertile abaxial surfacewith 34 bisporangiate synangia per lobe.Ovulate strobili usually 12, erect, cylindrical

to ovoid, up to 9 cm long, 4.5 cm in diam, brownto reddish tomentulose when young, dark-greenand glabrous when mature, apex acute; peduncle

brown tomentose, 3 cm long, 1.1 cm in diam;megasporangiophores peltate, distal face hexag-onal-truncate when immature, scutiform whenmature 0.81.2 cm high, 1.31.9 cm wide. Seedovoid, 1.6 cm long, 1.3 cm in diam, sarcotesta

pink when young and orange at maturity.Chromosome number.2n=16 (Marchant,

1968; Moretti et al., 1991; Stevenson et al.,19951996a).

Distribution and habitat.Zamia scheriis endemic to Mexico (Tamaulipas, San LuisPotos, Quertaro and Hidalgo; Fig. 5), be-tween 140900 m elevation. The vegetationtype of its habitat is pine-oak forest, tropicaldeciduous forest and mountain tropical forest(sensu Rzedowski,1978).

Etymology.The specic epithet is inhonor of Friedrich Fischer, a German cycad

horticulturist of the 19th

century.Distinguishing features.Zamia scheriis

a small fern-like plant (up to 100 cm tall),

FIG. 5. Distribution ofZamia cremnophila ( ),Z. furfuracea ( ),Z. inermis (),Z. katzeriana ( ),Z. lacandona

(

),Z. monticola( ),Z.onanreyesii( ),Z. prasina( ),Z. sandovallii(

),Z. soconuscensis( ),Z. vazquezii ( ).



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with papyraceous but serrulate leaets andunarmed petiole and rachis. Ovulate strobiligreenish upon maturity.

Additional specimens examined.MEXICO.QUERTARO: Jalopan, Lpez 438 (XAL), Servn 1471(XAL), Vovides 330 (XAL). SAN LUIS POTOS: O. M.Clark 6839 (MO), J. Rees 1686(XAL), Stevenson et al.566 (MEXU, NY), Vovides 753 (XAL); El Naranjo, F.

Nicolalde-Morejn & J. Gonzlez Astorga 1614 (XAL),1615 (XAL), 1616 (XAL), 1617 (XAL), 1618 (MEXU,XAL), 1619(XAL); Ciudad Valles,F. Nicolalde-Morejn& J. Gonzlez-Astorga 1620 (XAL), 1621(XAL), 1622(XAL), 1623(MEXU, XAL), 1624(MEXU, XAL), 1625(MEXU, XAL), 1626 (MEXU, XAL). HIDALGO: Pisa-ores, O. Alcntara-Ayala & R. Mayorga-Saucedo 3325(FCME).

Zamia furfuracea L. f. in Aiton, HortusKew. 3: 477. 1789. Type: Palma america-na crassis rigidisque foliis, pl. 210, inHerm. Paradisus Batavus. 1698 (lectotype,designated by Stevenson & Sabato, 1986:

pl. 210).

Zamia muricatavar.obtusifoliaMiquel, Tijdschr. nat.Gesch. Physiol. 10(1): 7172. 1843. Zamia latifo-lia Loddiges ex Miquel, Tijdschr. wis-en natuurk.Wet. 2(4): 298. 1849. Type: Tab. VII, g. a inLinnaea 19(4): 1847. (neotype, designated byStevenson & Sabato,1986: Tab. VII, g. a).

Zamia furfuracea var. trewii A. DC., Prodr. 16(2):541. 1868. Type: Palmifolia fructu clavato poly-

pireno[polyspermo]. C. J. Trew, PI. Select. Tab.26. 1752 (holotype: Tab. 26, G; typotype, desig-nated by Stevenson & Sabato,1986: BM).

Stem hypogeous, becoming epigeous withage, often dichotomously branched, up to60 cm long, 20 cm in diam. Cataphyllscoriaceous, persistent, base triangular, apexlong aristate, up to 1034.5 cm at base,yellowish tomentose. Ptyxis inexed. Leaves3 to many, 451901030 cm, diffuse,

brown-yellowish when emerging; petiole1750 cm long, brown-yellowish whenyoung, subterete, armed with small pricklesup to 3 mm long; rachis terete, up to 120 cmlong, with few prickles along the proximalthird.Leaets818 pairs, sessile, coriaceous,obovate to oblanceolate, opposite to subop-

posite, imbricate, keeled, apex rounded tosub-acute, base attenuate, margins slightlyserrulate along the 1/2 distal portion, sub-

revolute; articulations yellow in young andjuvenile leaets, 0.40.7 cm wide; medianleaets 142047.5 cm. Pollen strobili

usually 24, erect, cylindrical, up to 17 cmlong, 1.8 cm in diam, yellowish to brown,apex acute; peduncle densely light-browntomentose, 12 cm long, 1.1 cm in diam;

pollen sporangiophores cuneiform, distal face

hexagonal-truncate, 0.450.55 cm long, fer-tile abaxial surface with 89 bisporangiatesynangia per lobe. Ovulate strobili usually 1

per crown, erect, cylindrical, up to 25 cmlong, up to 10.5 cm in diam, yellowish-greentomentulose when immature, light brownupon maturity, apex apiculate; peduncle yel-lowish tomentose, up to 8 cm long, up to1.3 cm in diam; megasporangiophores pel-tate, distal end hexagonal-truncate, 0.9 cmhigh, 1.3 cm wide. Seeds ovoid, sarcotesta

yellowish-green when immature turning redat maturity, 1.6 cm long, 1.2 cm in diam,sclerotesta smooth.

Chromosome number.2n= 18 (Moretti,1990a,b)


percentage of polymorphic loci is P= 90.7, theexpected heterozygosity is HE=0.356 andthe genetic differentiation between the two

populations currently under study is Fst=

0.161 (Gonzlez-Astorga et al. unpubl. data).Distribution and habitat.Endemic toMexico in central and south coastal Veracruz,along a coastal stretch of approximately150 kmin stable dunes and basalt cliffs (Fig.5).

Etymology.The specic epithet alludes tothe persistent brown-yellowish trichomes ofthe leaves throughout the developmentalstages of the plant.

Distinguishing features.Leaves stronglykeeled with highly imbricate and coriaceousobovate to oblanceolate leaets with brown-yellowish indumentum persisting with age.

Additional specimens examined.MEXICO.VERACRUZ: Ibarra-Mariquez 316 (MEXU, MO),

Ibarra-Mariquez 1952, 1959 (MEXU); Alvarado, Rees1650, 1651, 1652(XAL),M. Vzquez-Torres et al. 4871(CIB); Catemaco,Calzada 1475(MEXU), Calzada 2451(XAL),Cedillo 2610(MEXU, XAL),Gonzlez-Quintero1520 (ENCB), Ibarra 316 (MEXU), Lot 1277-14 (F,XAL), Menendez 115(MEXU, MO, XAL),F. Nicolalde-

Morejn et al. 1484 (XAL), 1485(XAL), 1486(XAL),J.Rees 1651(IBUG, MEXU, XAL), J. Rees 1652 (XAL),

Schatz & Nee 207(XAL), Vovides 567 (MEXU, XAL);Mecayapan, Calzada et al. 11325 (XAL), Castillo-Campos 12732 (XAL); Lerdo De Tejada, Vovides 828(XAL), 829 (XAL), 830 (XAL), 831 (XAL), 832 (XAL),



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833 (XAL), 839 (XAL); San Andrs Tuxtla, Castillo-Campos et al. 13881 (XAL), Chazaro B. 512 (XAL),

Hammel & Merello 15499 (MO), Hernndez-M 1216A(F, MEXU), Lorence 4978 (MEXU), Sousa 3099 (F,MEXU, MO),Vovides & Iglesias 1148 (XAL).

Zamia herrerae Caldern & Standl., Proc.Wash. Acad. Sci. 14(4): 93. 1924. Type: ElSalvador. Vicinity of Sonsonate, 17 Jul1923,S. Caldern 1682 (holotype: US).

Stem hypogeous, bifurcate in adult plants,6264.513.5 cm in diam.Cataphyllschar-taceous, persistent, base triangular, apexaristate, 4.61.2 cm wide at base, yellowishtomentose.Ptyxisinexed.Leaves24, erect,

green to light-brown when emerging, 61

9624.529.5 cm; petiole 1934 cm long, brown-ish in young leaves, terete, armed with

prickles up to 3 mm long; rachis subterete,up to 63 cm long, with few prickles along the

proximal third. Leaets 1532 pairs, sessile,papyraceous, lanceolate, alternate to subop-posite, apex acute, base symmetric attenuate;margins dentate along distal 2/3, subrevolute;articulations dark brown when young, 0.30.5 cm wide; median leaets 22383.1

4.4 cm. Pollen strobili usually 2

3, erect,cylindrical to conical, 4.37.5 cm long, 1.32.1 cm in diam, light brown tomentulose,apex mucronate; peduncle densely light-

brown tomentose, 6.27.8 cm long, 1.11.4 cm in diam; pollen sporangiophorescuneiform, distal face hexagonal, 0.35 cmlong, fertile abaxial surface with 34 bispor-angiate synangia per lobe. Ovulate strobiliusually solitary, erect, cylindrical to ovoid,7.111.6 cm long, 4.14.9 cm in diam,

brown, tomentulose, apex acute; peduncledensely brown tomentose, 3.54.2 cm long,1.11.3 cm in diam; megasporangiophores

peltate, distal face hexagonal-truncate, 1.31.6 cm high, 2.22.8 cm wide. Seeds ovoid,sarcotesta pink when immature, red at matu-rity, 1.61.9 cm long, 1.21.5 cm in diam,sclerotesta smooth.

Chromosome number.2n=23, 24 (Fig.4).Distribution and habitat.The species

range is El Salvador, Guatemala, and southernMexico in Chiapas, between 100600 m

elevations (Fig. 6). It generally grows intropical deciduous forest (sensu Rzedowski,1978) on deep clay soils. Also, this cycad may

be found in secondary growth forests andpastures.

Etymology.The specic epithet honorsHector Herrera, a scientist from El Salvador.

Distinguishing features.Zamia herreraeis

distinguished by its long-lanceolate papyra-ceous leaets with dentate margins along thedistal 2/3.

Additional specimens examined.MEXICO.CHIAPAS: Escuintla, Matuda 16368 (MEXU), 16871(MEXU), 17332 (MEXU), 18332 (MEXU), M. A.

Prez-Farrera 143 (HEM), Schutzman 526(XAL), 527(XAL), 528 (XAL); Tonal, Farrera 2489 (CHIP), F.

Nicolalde-Morejn & J. Gonzlez-Astorga 1579 (XAL),1580 (XAL), 1581 (XAL), M. A. Prez-Farrera 744(CIB).

GUATEMALA. P. C. Standley 67306(F).NICARAGUA. MANAGUA: Cultivated, A. Grijalva3658(MO).

Zamia inermis Vovides, J.D. Rees & Vzq.Torres, Flora de Veracruz 26: 22. 1983. Type:Mexico. Veracruz: 6 Jun 1981, Vovides 666(holotype: XAL; isotype: F). (Fig.8)

Stemepigeal, erect, dichotomously branch-ing in mature plants, 1543 cm long, 8.6

26.4 cm in diam. Cataphylls chartaceous,persistent, base triangular, apex aristate, 2.12.64.15.6 cm wide at base, yellowishtomentose.Ptyxisinexed to erect.Leaves1035, erect, light to yellowish-green when emerg-ing, 309543.560 cm; petiole 1841 cm long,greenish in young leaves, subterete, without

prickles; rachis subterete, 1519 cm long,unarmed. Leaets 2732 pairs, sessile, coria-ceous, linear-lanceolate, opposite to suboppo-site, apex acute, base attenuate; margins entire,subrevolute; articulations 0.40.6 cm wide;median leaets 2030.50.91.2 cm. Pollen

strobilicylindrical, usually 12 per crown, erect,up to 9.1 cm long, up to 2.8 cm in diam, beige-yellowish, apex acute; peduncle densely light-yellowish tomentose, up to 4.5 cm long, 1.1 cmin diam; pollen sporangiophores cuneiform,distal face hexagonal-truncate, 0.35 cm long,fertile abaxial surface with 56 bisporangiatesynangia per lobe.Ovulate strobili usually 12

per crown, erect, cylindrical, 1323 cm long, 89.8 cm in diam, light-brown to beige tomentu-

lose, apex apiculate; peduncle brown tomentose,68 cm long, 1.21.4 cm in diam; megaspor-angiophores peltate, distal face hexagonal-trun-



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cate, 0.70.9 cm high, 1.11.3 cm wide.Seedsovoid, sarcotesta pink when immature, red atmaturity, 1.72.5 cm long, 1.42.1 cm in diam,sclerotesta smooth.

Chromosome number.2n= 16 (Vovides,1983).

Distribution and habitat.Endemic toMexico in a small mountain range in centralVeracruz (Fig. 5) at 150300 elevation on

basaltic soils. The vegetation type of the habitatis tropical deciduous forest (sensuRzedowski,1978).

Etymology.The specic epithet alludes tothe absence of prickles on the petiole andrachis and the entire leaet margins.

Distinguishing features.Zamia inermisdiffers from its congeners in Mexico by atotal absence of prickles along the petiole andrachis, as well as having totally entire leaetsand light-yellowish to clear-beige tomentu-lum on the pollen and ovulate strobili.

Additional specimens examined.MEXICO.VERACRUZ: Actopan, Acosta & Acosta 234 (XAL), F.

Nicolalde-Morejn & Vovides 1415 (XAL), 1416(XAL),1417 (XAL), Schutzman 570 (XAL), 571 (XAL), 572(XAL), 575 (XAL), 576(XAL), 577(XAL), J. Rees etal. 681(XAL).

Zamia katzeriana (Regel) Rettig, Garten-ora 45: 148. 1896. Ceratozamia katzeri-ana Regel, Acta Horti Petrop. 4(4): 298.1876. Type: ex Horto Katzer., Regel s.n.(lectotype, designated by Stevenson &

Sabato,1986: LE).

Zamia splendens Schutzman, Phytologia 55(5): 299.1984. Type: Cultivated in Fairchild Tropical Garden,Miami, accession no. FTG 761046, 11 Apr 1984, J.Watson s.n. (holotype: NY; isotypes: FLAS, FTG,MEXU).

Stemhypogeous, unbranched, up to 25 cmlong, up to 7 cm in diam. Cataphylls charta-ceous, semidecidious, base triangular, apexaristate, 5.3 1.4 cm at base, yellowish

tomentose. Ptyxis inexed. Leaves 12 (3),492203558 cm, ascending to descending,

bright reddish-pink with lustrous cuticle when

FIG. 6. Distribution ofZamia scheri (), Z. herrerae ( ), Z. loddigesii ( ), Z. oreillyi ( ), Z. paucijuga ( ), Z.

polymorpha(),Z. purpurea (),Z. spartea( ),Z. standleyi( ),Z. tuerckheimii ( ),Z. variegata( ).



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emerging, dark green when mature; petiole20130 cm long, terete, armed with fewsimple prickles; rachis subterete, up to84 cm long, unarmed. Leaets 37 pairssessile, coriaceous, oblong-lanceolate, oppo-site to subopposite, adaxial surface with

brilliantly shining cuticle throughout life ofleaet, apex acute, base attenuate; marginsdentate along distal third, subrevolute; artic-ulations brown in young leaets turning greenwith age 0.61.6 cm wide; median leaets18353.512 cm. Pollen strobili 15, con-ical, light-brown tomentulose, erect uponemergence becoming prostrate to decumbentupon maturity, up to 3.9 cm long, 1.1 cm indiam, apex acute; peduncle light-brown to-

mentose, up to 5.8 cm long, 1.2 cm in diam;pollen sporangiophores cuneiform, distal facehexagonal scutiform, 0.35 cm long, fertile

abaxial surface with 56 bisporangiate syn-angia per lobe. Ovulate strobili usuallysolitary, decumbent to erect, elliptic, 812 cm long, 4.56 cm in diam, brown toyellowish tomentose, apex aristate; peduncle

brown tomentose, 2.14.3 cm long, 1.11.3 cmin diam; megasporangiophores peltate, distalface hexagonal-truncate to scutiform, 11.2 cmhigh, 1.11.3 cm wide.Seedsovoid, sarcotesta

pink when immature, red at maturity, 1.11.4 cm long, 1.61.8 cm in diam, sclerotestasmooth.

Chromosome number.2n=16 (Schutzman,1984; Moretti,1990a).

Diversity and genetic structure.Theaverage of alleles per locus is A=1.95, the

percentage of polymorphic loci is P= 84.4,the expected heterozygosity is HE=0.280and the genetic differentiation between

FIG. 7. Zamia scheriA. Habit.B. Leaets. (FromStevenson 566, NY).



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FIG. 8. Zamia inermis. AB. Habit. C. Cataphyll. D. Pollen strobilus. E. Microsporophyll, abaxial and adaxialview.FH. Ovulate strobilus. I. Ovulate sporangiophores. J. Seed. (From the holotype.)



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the two populations currently under study isFst=0.194 (Gonzlez-Astorga et al. unpubl.data).

Distribution and habitat.Endemic toMexico and known from the states of

Chiapas, Tabasco and Veracruz (Fig. 5) at200700 m in evergreen tropical forest (sensuRzedowski,1978).

Etymology.Specic epithet in honor ofKatzer, inspector of the gardens in Paullowsk(Stevenson & Sabato,1986).

Distinguishing features.Zamia katzerianais easily distinguished from its congeners byhaving leaves with highly lustrous or shinycuticles. Emerging leaves are a bright reddish-

pink. Both pollen and ovulate strobili are

borne on long peduncles that become descen-dent to prostrate.

Additional specimens examined.MEXICO.VERACRUZ: Las Choapas, Martnez & Martnez-M. 825(HEM), F. Nicolalde-Morejn et al.1436 (XAL), 1437(XAL).TABASCO: Teapa, Hdez-Najarro 622 (CHIP), M.

A. Magaa 1905 (MEXU), M. A. Prez-Farrera s.n.(XAL), M. A. Prez-Farrera 899 (HEM, MEXU),Walters s.n. (FTG accession 12-2, XAL). CHIAPAS:SanFernando, Vovides et al. 1266 (XAL), Palacios 383(CHIP), F. Nicolalde-Morejn & Prez-Farrera 1420(XAL), M. A. Prez-Farrera s.n. (XAL), Walters s.n.

(FTG accession 23-2, XAL); Ocozocoautla, Gmez-Pompa 705 (FCME, MEXU), F. Nicolalde-Morejn etal. 1453(XAL), 1454 (XAL), 1455 (XAL), 1456(XAL),1457(XAL), 1458 (XAL), 1459 (XAL), 1460 (XAL),M.

A. Prez-Farrera 29 (CHIP, CIB, MEXU); Tila, Vovideset al. 1340 (XAL), 1343(XAL), 1341(XAL).

Zamia lacandona Schutzman & Vovides,Novon 8(4): 441. 1998. Type: Mexico.Chiapas: Selva Lacandona, July 1984,Schutzman 517 (holotype: FLAS; isotype:XAL).

Stem hypogeous, unbranching, 614 cmlong, 59 cm in diam.Cataphyllscoriaceous,

persistent, base triangular, apex long-aristate,8.33.6 cm at base, reddish-brown tomen-tose. Ptyxis inexed, reddish-brown. Leavesusually solitary, up to 3 under cultivation, 461713275 cm, ascending, reddish-brownwhen emerging; petiole 1495.3 cm longwith bulbous base, blackish in young leaves,subterete, proximal section strongly chan-

neled, armed with prickles up to 5 mm long;rachis subterete, 3176 cm long, with few

prickles along proximal third. Leaets 417

pairs, sessile, coriaceous, lanceolate, oppositeto subopposite, subfalcate, apex acute, baseattenuate, margins dentate along distal third,subrevolute; articulations dark-brown whenyoung, 0.41.1 cm wide; median leaets

15.6372.96 cm. Pollen strobili usually23, erect, conical, 5.46.6 cm long, 1.51.7 cm in diam, light-brown, apex acute;

peduncle light-brown tomentose, 5.86.9 cmlong, 1.2 cm in diam; pollen sporangiophorescuneiform, distal face hexagonal truncate,0.35 cm long, fertile abaxial surface with 45 bisporangiate synangia per lobe. Ovulate

strobili usually solitary, erect, ellipsoid,13.2 cm long, 4.9 cm in diam, dark-brown,tomentulose, apex acute to slightly apiculate;

peduncle densely brown tomentose, 6.5 cmlong, 1.2 cm in diam; megasporangiophorespeltate, distal face hexagonal-truncate, 11.2 cm high, 11.5 cm wide.Seedsirregularlyovoid, sarcotesta pink when immature, red atmaturity, 22.4 cm long, 1.31.9 cm in diam,sclerotesta smooth.

Chromosome number.2n=16, 17, 18(Schutzman & Vovides,1998).


percentage of polymorphic loci is P=67.9, theexpected heterozygosity is HE=0.191 andthe genetic differentiation between the two


Distribution and habitat.Endemic toChiapas (Fig. 5) in the vicinity of theLacandon forest at 80200 m elevation, inevergreen tropical forest (sensu Rzedowski,1978). It is also found in secondary succes-sion stages of the tropical forest.

Etymology.The epithet is derived fromthe name of the 1.8 million hectare SelvaLacandona (Lacandona Jungle) in southeast-ern Chiapas, which itself bears the name ofthe Lacandona Maya Indians who inhabit theforest.

Distinguishing features.This species isdistinguished by having a solitary large leaf(up to three leaves may be maintained on

plants under cultivation) with a stout erectpetiole and a strongly bulbous base. The leafis reddish-brown at emergence and petiole

dark purplish-brown turning dark-brown withage. Ovulate cone generally solitary, withacute apex.



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Additional specimens examined.MEXICO.CHIAPAS:Palenque, Schutzman 510 (XAL), 511 (XAL),512 (XAL), 513 (XAL), 514 (XAL), 515 (XAL), 516(XAL), 517(XAL), 518 (XAL), 519(XAL), 520 (XAL),

F. Nicolalde-Morejn & N. Martnez 1418 (XAL),M. A.Prez-Farrera 890 (HEM, MEXU), Walters s.n. (FTG

accession 14-2, XAL), M. Vzquez-Torres et al. 3925(CIB); San Jernimo Tulija, Chavelas et al. ES=315(ENCB, MEXU), Schutzman 521 (XAL), 522 (XAL),524(XAL), 523(XAL), 525(XAL).

Zamia loddigesii Miq., Tijdschr. Natuurl.Gesch. Physiol. 10: 72. 1843. Type: culti-vated byVan Houtte 3374 (lectotype,heredesignated:U).

Zamia galeotti De Vriese, in Hoven & De Vries.

Tijdschr. Natuurl. Gesch. Physiol. 12: 24. 1845.Type: Mexico. Veracruz: 5 July 1983. D. W.Stevenson 538 (neotype, here designated: NY,isoneotype: XAL).

Zamia leiboldii Miq., Linnaea 19: 425. 1847. Zamialoddigesiivar.leiboldii(Miquel) A. DC., Prodr. 16(2): 541. 1868. Type: E. Mexico in HortumLoehrianum Lipsiae attulit Liebold, 1845, Miquel

s.n. (holotype: U).Zamia loddigesii var. angustifolia Regel, Bull. Soc.

Nat. Moscou 30(1): 190. 1857. Type: ex hortoPetropolitano, 1856,Regel s.n.(holotype: LE)

Zamia loddigesii var. obtusifolia Regel, Bull. Soc.

Nat. Moscou 30(1): 190. 1857. Type: t. 186,

gs2728 in Gartenora 6: 1857. (lectotype, designat-ed by Stevenson & Sabato,1986: LE).

Zamia mexicanaMiquel, Prodr. syst. Cycad. 13. 1861.Type:Eriozamia mexicanaH. Belg., 1847, Miquel

s.n. (holotype: U).

Zamia leiboldii var. angustifolia Regel, Trudy Imp.S.-Petersburgsk. Bot. Sada 4(4): 307. 1876. Type:Mexico. Oaxaca: 15 Jul 1983.D. W. Stevenson 559(neotype,here designated:NY; isoneotype, XAL).

Zamia leiboldii var. latifolia Regel, Trudy Imp. S.Petersburgsk. Bot. Sada 4(4): 307. 1876. Type: exHorto Petropolitano, 1875, Regel s.n. (holotype:LE).

Zamia lawsoniana Dyer in Hemsley, Biol. Centr.-Amer., Bot. 3(16): 195. 1884. Type: Mexico.Oaxaca:Fielding 209(holotype: OX; isotype: K)

Zamia cycadifolia Dyer in Hemsley, Biol. Cent.-Amer., Bot. 3(16): 195. 1884, non Jacquin 1809.Zamia loddigesii var. cycadifolia Schuster, Pan-zenreich 99: 148. 1932. Type: Mexico. Bourgeau s.n. (holotype: K; isotype: C). nomen illegit.

Zamia sylvatica Chamberlain, Bot. Gaz. 81: 223.1926. Type: Mexico, Oaxaca, Tuxtepec, Sep 1910,C. J. Chamberlain s.n. (lectotype, designated byStevenson & Sabato, 1986: NY; isolectotype, F-3sheets).

Zamia loddigesii var. angustifolia (Regel) J. Schust.,in Engl., Panzenr. 4(1): 148. 1932. Type: Mxico.Veracruz: savanne bei Mundo nuevo, Karwinski

1028b (lectotype, designated by Stevenson &Sabato,1986: LE; isolectotype, LE).

Zamia loddigesii var. longifolia J. Schust., in Engl.,Panzenr. 4(1): 147. 1932. Type: Mexico. Vera-cruz: Colipa,Karwinski 1029 (lectotype, designat-ed by Stevenson & Sabato,1986: LE; isolectotype:

LE).

Stemhypogeous, branching dichotomouslywith age, 1045 cm long, 815 cm in diam.Cataphyllschartaceous, persistent, base trian-gular, apex aristate, 8.4 3.7 cm at base,yellowish tomentose. Ptyxis inexed. Leaves23 (4) ascending to spreading, 45963041 cm, light-green when emerging, green todark-green when mature; petiole 1525 cmlong, green in young leaves, subterete, armed

with prickles up to 4 mm long; rachissubterete, up to 57 cm long, with few prickleson the proximal third. Leaets 1223 pairs,sessile, coriaceous, linear-lanceolate, oppositeto subopposite, apex acute, base attenuate,margins serrulate along distal third, subrevo-lute; articulations 0.40.7 cm wide; medianleaets 16261.83.1 cm.Pollen strobili12 per crown, up to 6 (7) when multiplecrowned, erect, cylindrical, 814 cm long,1.83.5 cm in diam, light brown tomentulose,

apex acute; peduncle light-brown tomentose,6 cm long, 1.2 cm in diam; pollen sporangio-phores cuneiform, distal face hexagonal-truncate, 0.3 cm long, fertile abaxial surfacewith 68 bisporangiate synangia per lobe.Ovulate strobiliusually 12 per crown, erect,ellipsoid to conical, up to 16 cm long, up to6 cm in diam, beige-tomentulose, apex acute;

peduncle brown-tomentose, up to 6 cm long,1.6 cm in diam; megasporangiophores peltate,distal face hexagonal-truncate, 0.71 cm high,1.92.6 cm wide.Seedsovoid, sarcotesta pinkwhen immature, red at maturity, 1.41.8 cmlong, 0.81 cm in diam, sclerotesta smooth.

Chromosome number.2n= 18 (Norstog,1980; Moretti,1990a,b).


percentage of polymorphic loci is P= 66.6,the expected heterozygosity is HE=0.266and the genetic differentiation between thetwo populations currently under study is

Fst=0.179 (Gonzlez-Astorga et al. unpubl.

data).Distribution and habitat.Endemic to

Mexico and distributed widely within the



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states of Tamaulipas, Hidalgo, Veracruz,Tabasco and parts of Oaxaca with a singleknown locality in Chiapas (Fig. 6). Thevegetation types are evergreen tropical forest,tropical deciduous and sub-deciduous forests

(sensu Rzedowski,1978), as well as a varietyof secondary succession and disturbed hab-itats such as pastures and cornelds, as wellas road-side vegetation.

Etymology.The specic epithet honorsConrad Loddiges (17381826), a Germanhorticulturist who lived in London andcultivated American cycads.

Nomenclatural notes.The descriptionof Zamia loddigesii was lectotypied byStevenson & Sabato (1986). However, it is

no longer permissible to designate a descrip-tion to serve as a type. Moreover, since thena specimen, Van Houtte 3374 at U, wasfound that was sent to Miquel by VanHoutte. This specimen predates the publica-tion of Z. loddigesii Miq. and matches thedescription. Thus, we are designating it thelectotype.

Distinguishing features.Zamia loddigesiiin contrast to Z. paucijuga, has been widelycollected along the Gulf of Mexico seaboard

and on the Yucatan peninsula since the 19thcentury, largely by British and Russianbotanists, and recently by Mexican and NorthAmerican researchers. The high morphologi-cal variation presented by this species hasresulted in the publication of 10 afne names(Hill et al., 2007) that at times, has been a

basis for the separation of natural popula-tions, i.e., that of the Yucatan peninsula(formerly Z. loddigesii sensu Vovides &Olivares,1996) that is now a separate entityZ. polymorpha (Stevenson et al., 19951996b), a decision based on both vegetativeand reproductive characters that differ fromZ.loddigesii mainly on pollen and ovulatestrobilus shape and indument color.

Zamia loddigesii is similar to Z. polymor-pha in leaf morphology; however, there aredifferences in the reproductive structures.Pollen cones of Z. loddigesii are beige incolor with a blunt apex whereas those ofZ.

polymorpha are dark reddish-brown or ma-roon with an acute apex. vulate cones ofZ.

loddigesii are cylindrical and beige in colorand those of Z. polymorpha are ovoid anddark maroon in color.

Additional specimens examined.MEXICO.CHIAPAS:Ocozocoautla, M. A. Prez-Farrera 81 (CHIP,HEM). HIDALGO: Atlapexco, San Juan 15 (XAL), 16(XAL), 17(XAL), 18 (XAL), 19 (XAL), 20 (XAL), 21(XAL).PUEBLA: Sarukn et al. 4632 (FCME, MEXU).Oaxaca, Tuxtepec,J. Chamberlain s.n. (MO).TAMAULI-PAS: R. L. Dressler 1858 (MO), Mayel et al. 791(MEXU); Aldama,F. Nicolalde-Morejn& J. Gonzlez-

Astorga 1585 (XAL), 1586(XAL), 1587 (XAL), 1588(XAL). VERACRUZ: Chavelas et al. ES-4231 (MEXU),

Dorantes et al. 964(MEXU, MO), Dorantes et al. 1112(MEXU), Lot 733 (MEXU), Medrano et al. 2725(MEXU), Nevling & Gmez-Pompa 140 (MEXU),Santos 353 (XAL, XALU); Acayucan, Vovides et al.1376(XAL), 1377(XAL); Actopan, J. I. Calzada et al.6369 (MEXU, XAL), Lot 1027(XAL), A. Vovides 754(XAL), 755 (XAL), 817(XAL), 818 (XAL), 819 (XAL),820 (XAL), 821 (XAL), 822 (XAL), 823 (XAL), 824(XAL); Alto Lucero,J. Rees 1627(XAL), 1629 (XAL),1630 (XAL), 1631 (XAL), 1632 (XAL), 1637 (XAL),Vovides 846 (XAL); Atoyac, Acevedo & Castillo-Campos 240 (XAL); Cotaxtla, Gonzlez 82 (MEXU);Chicontepec, J. Rees 1615 (MEXU, XAL); Choapas,Vovides et al. 1373, 1374, 1735 (XAL); Coatepec, J.

Rees & Vovides 1670(XAL); Coatzacoalcos,Castillo &Acosta 16220(XAL); Colipa,J. Rees 1634, 1635(XAL);Cosautlan,Vovides 35 (XAL); Emiliano Zapata, J. Rees1763 (XAL), Stevenson et al. 538 (MEXU, NY), M.Vzquez-Torres 8071 (CIB); Huejultla, Stresser 291(MEXU); Hueyapan de Ocampo, Gmez-Pompa 4424(XAL), Vovides et al. 1378 (XAL), 1379 (XAL), 1380(XAL); Jalcomulco,Castillo & Zamora C. 7542(XAL),Castillo & Gmez-Pompa 2588(XAL),Castillo-Campos

2727 (XAL); Mecayapan, Castillo et al. 13681 (XAL),13792(XAL), 13843(XAL), 13861(XAL), 13865(XAL),13866 (XAL), A. Calatayud & J. Martnez-Gndara 124(CIB); Moloacan, J. Rees 1656 (XAL); Puente Nacional,Castillo & Medina 4261(XAL); Papantla,R. Cuevas et al.4652(ZEA); Soteapan,Leonati 42(MEXU); Soteapan, A.Calatayud & J. Bentez R 285 (CIB),M. A. Santos R. 352(CIB); Tampico Alto, Ortega & Ortega O. 2437 (XAL);Tezonapa,Robles 370(XAL); Totutla,J. Rees 1661(XAL);Yecuatla,J. Rees 1633(XAL).

Zamia monticola Chamb., Bot. Gaz. 81:219. 1926. Type: cultivated from a single

seed collected opposite the crater of Nao-linco, near Xalapa, Veracruz, Mexico, Oct1925, C. J. Chamberlain s.n. (holotype:MO; isotype: NY).

Stemepigeal, up to 30 cm tall, 1820 cm indiam.Cataphyllsbase triangular, apex linear-lanceolate, 3612 cm wide at base.Leaves520, 100200 cm long, erect to slightlycurved; petiole 5075 cm long, terete, armedwith stout prickles in lower half; rachis terete,

up to 100 cm long, with few prickles alongthe lower half. Leaets 3040 pairs, sessile,chartaceous to papyraceous, linear-lanceolate,



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opposite to subopposite, subfalcate near thebase, apex long-acuminate and often stronglycurved, base attenuate, margins serrulate onlynear the base, subrevolute; articulations0.40.7 cm wide; the median leaets 25

3046 cm. Pollen strobili usually 26,erect, cylindrical to oblong, 1220 cm longand 24 cm in diam, cream to light brown,apex acute; peduncle light brown tomentose,1020 cm long; pollen sporangiophores cu-neiform, distal face hexagonal, 0.4 cm long,fertile abaxial surface 2-lobed with 1016

bisporangiate synangia per lobe. Ovulatestrobiliunknown.

Chromosome number.UnknownEtymology.The specic epithet alludes to

its mountainous habitat type, originallythought to be near Naolinco near the city ofXalapa, Veracruz in Mexico. It is now knownthat this species is not known from Mexicoand is endemic to Guatemala.

Distribution and habitat.Endemic toGuatemala on rocky outcrops in primary andsecondary evergreen tropical rainforest (Fig.6).

Distinguishing features.Characterized byits consistently chartaceous to papyraceouslong acuminate leaets that are strongly

curved near the apex with light serrulations.Additional specimens examined.GUATEMALA.

ALTA VERAPAZ:H. Frther 2621/592(NY).

Zamia onanreyesii C. Nelson & G. San-doval. Ceiba 49(1): 135. 2008. Type:Honduras. Corts, 7 Jan 2008, O. Reyes406(holotype: TEFH).

Zamia bussellii Schutzman, R. S. Adams, J. L.Haynes & Whitelock. The Cycad Newsletter 31

(2/3), 22. 2008. Type: Honduras. Corts, June2003,Whittington 2003/01(holotype: FLAS).

Stemup to 2 m tall, up to 16 cm in diam.Ptyxis inexed. Leaves 315(44), 601801550 cm, erect to slightly curved, tomentu-lose when emerging; petiole 1540 cm long,terete, sparsely to moderately armed with

prickles; rachis terete, up to 40120 cm long,with few prickles along proximal third. Leaf-lets up to 30 pairs, sessile, subcoriaceous,oblong-lanceolate, opposite to subopposite,

long acuminate apically, base attenuate, mar-gins serrulate along distal third, subrevolute;articulations yellowish; median leaets to

364 cm. Pollen strobili usually 13, erect,cylindrical, up to 27.5 cm long, up to 4 cm indiam, light brown to tan, apex acute; pedun-cle brown to tan, tomentose, up to 8.5 cmlong, up to 1.6 cm in diam; pollen sporan-

giophore cuneiform, distal face hexagonaltruncate, fertile abaxial surface with up to 9

bisporangiate sori per lobe. Ovulate strobiliusually solitary, erect, cylindrical, up to43 cm long, 12 cm in diam, brown togreenish, tomentulose, apex conical; peduncledensely brown tomentose, up to 5 cm long,up to 2.5 cm in diam, distal face hexagonal-truncate, 2.9 cm high, 4.7 cm wide. Seedsovoid, sarcotesta red at maturity, up to 3 cmlong, up to 2 cm in diam, sclerotesta smooth.

Chromosome number.

Unknown.Distribution and habitat.The speciesrange is Honduras (Fig.6), between 01300 melevations in evergreen tropical forest.

Etymology.The specic epithet is inhonor of Onn Reyes, a Honduran biologist.

Distinguishing features.Stems arbores-cent up to 2 m tall, leaets coriaceous, longacuminate apically, with margin serrulate.

Nomenclatural note.BothZ. onanreyesiiandZ. bussellliiwere published in 2008. The

dates of issue are however different. The dateof issue for Z. onanreyesii is 6 September2008 and the date of issue forZ. bussellii is16 October 2008. Thus, under Article 29.1 ofthe Interantional Code of Botanical Nomen-clature (McNeill et al., 2007), Z. onanreyesiihas priority and is used here.

Additional specimen examined.HONDURAS.Departamento Corts: J. Haynes et al. 044A, 044B(TEFH).

Zamia oreillyiC. Nelson, Ceiba 46(12): 56.2005. Type: Honduras. Atlntida: 8 Apr2006, G. Sandoval et al. 1157 (holotype:TEFH). (Fig.9)

Stem hypogeous, non-branching, up to25 cm long, up to 7.5 cm in diam. Leavesusually 1(2), up to 78.2 cm long, 3135 cmwide, ascending to descending; petiole up to47 cm long, subterete, armed with small

prickles; rachis subterete, up to 3540 cmlong, with few prickles along the proximal

third. Leaets 2931 pairs, sessile, papyra-ceous to sub-coriaceous, linear-lanceolate,opposite to subopposite, imbricate, apex



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acuminate, base attenuate, margins dentate torarely entire along distal third, up to 0.3 cm,subrevolute; articulations brown in youngleaets, 0.30.5 cm wide; the median leaets

up to 16 cm long, 1 cm wide.Pollen strobiliusually solitary, decumbent, cylindrical, up to2.5 cm long and 1 cm in diam, light brown,

apex acuminate; peduncle light brown tomen-tose, up to 15.9 cm long, 0.2 cm in diam;

pollen sporangiophores cuneiform, distal facetruncate-hexagonal, 0.4 cm long, fertile ab-

axial surface, with 23 bisporangiate synan-gia per lobe. Ovulate strobiliunknown.

Chromosome number.Unknown

FIG. 9. Zamia oreillyi. A. Habit. B. Cataphyll. CE. Leaet variability. C. Broad and imbricate with dentatemargins.D. Narrow and imbricate with dentate margins. E. Narrow and slighly imbricate with entire margins. F.Ovulate strobilus. G.Pollen strobilus.H. Microsporophyll, abaxial view. (A-E, G-H fromBalick 1711, NY).



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Etymology.The specic epithet is inhonor of Carlos Manuel OReilly, a Hon-duran biologist.

Distribution and habitat.Endemic toHonduras, between 0200 m in evergreen

tropical rainforest (Fig.6).Distinguishing features.Characterized by

strongly imbricate linear-lanceolate tooblong leaets with dentate margins alongdistal third; pollen strobili usually solitary,decumbent, with fertile section up to 2.9 cmlong and decumbent peduncle up to 15.9 cmlong.

Additional specimen examined.HONDURAS.ATLNTIDA.Balick 1711(NY, TEFH).

Zamia paucijuga Wieland, American FossilCycads 2: 212. 1916. Type: Fig. 86 inAmerican Fossil Cycads 2: 212.1916.(lectotype, designated by Stevenson &Sabato,1986: Fig. 86).

Stemhypogeous, branching dichotomouslywith age, 1527 cm long, 813 cm in diam.Cataphyllscoriaceous, persistent, base trian-gular, apex aristate, 4.5 3.4 cm at base,

brown tomentose.Ptyxis in

exed.Leaves 2

3, ascending to descending 41952936 cmwide, brownish when emerging; petiole 10.232 cm long, green in young leaves, subterete,armed with prickles up to 4 mm long; rachissubterete, up to 56 cm long, with few pricklesalong the proximal third. Leaets 528 pairssessile, coriaceous, lanceolate, opposite tosubopposite, apex acute, base attenuate, mar-gins serrulate to slightly dentate along distalthird, subrevolute; articulations brownish inyoung leaets, 0.40.6 cm wide; the medianleaets 14192.33.4 cm. Pollen strobiliusually 12, erect, cylindrical, 6.311 cmlong, 2.12.6 cm in diam, light browntomentulose, apex acute; peduncle light-

brown tomentose, 6.3 cm long, 1.2 cm indiam; pollen sporangiophores cuneiform, dis-tal face hexagonal-truncate, 0.3 cm long,fertile abaxial surface with 68 bisporangiatesynangia per lobe. Ovulate strobili usuallysolitary, erect, ellipsoid to cylindrical, 8.1 cmlong, 5.2 cm in diam, brown-yellowish,

tomentulose, apex apiculate; peduncle dense-ly brown tomentose, 3.9 cm long, 1.3 cm indiam; megasporangiophores peltate, distal

face hexagonal-truncate, 0.60.8 cm high,1.11.5 cm wide. Seeds ovoid, sarcotesta

pink when immature, orange at maturity, 22.8 cm long, 1.51.7 cm in diam, sclerotestasmooth.

Chromosome number.2n= 23, 34, 25, 26,27, 28 (Moretti & Sabato,1984).

Distribution and habitat.Endemic toMexico. Known from the states of Nayarit,Jalisco, Colima, Michoacan, Guerrero, andOaxaca (Fig. 6). Found in varied habitatsfrom dry open woodlands to understoryevergreen tropical forest.

Etymology.The specic epithet alludes tofew leaet pairs per leaf.

Distinguishing features.Zamia paucijuga

is highly variable morphologically and inchromosome number and karyotype (Moretti,1990a,b). This may represent more than oneentity under the concept of Z. paucijuga

sensu stricto Wieland (1916). This speciesshares a number of characteristics with Z.loddigesii, being small plants with under-ground contractile stems and having coriaceousleaets with marginal teeth. Nevertheless, itdiffers from Z. loddigesii in having highlycoriaceous leaets with longer marginal teeth,

and seeds with an orange sarcotesta at maturityas contrasted to the red sarcotesta of Z.paucijuga. Zamia paucijugaoccurs along thePacic seaboard of Mexico whereas Z. loddi-

gesii is known generally from the Gulf ofMexico seaboard.

Additional specimens examined.MEXICO.COLIMA: McVaugh 15768(FCME, MEXU).GUERRERO:Acapulco, N. Noriega-Acosta 463, 546 (FCME), W.Thomas & J. L. Contreras 3744 (FCME); Chilpancingo,

Kruse 902 (FCME, MEXU), R. M. Fonseca 1210(FCME), F. Nicolalde-Morejn et al. 1566, 1567, 1568,

1569 (XAL), Jos Azueta, Vovides et al. 1426 (XAL);Petatln, Vovides et al. 1427, 1428, 1429, 1430, 1431,1432, 1433, 1434 (XAL); Unin de Isidro Montes DeOca,Vovides et al. 1416, 1417, 1418, 1420, 1421(XAL);La Unin, G. Lozano-Valdez 331 (FCME), J. Jimnez331(FCME).JALISCO:Gmez-Pompa 4876(MEXU); ElArenal, Castillo et al. 9822 (XAL); Cabo Corrientes,Castillo et al. 10147, 11733, 10280, 10466 (XAL), J.Ceja et al 1437(UAMIZ),J. Ceja et al. 1470 (UAMIZ),

F. Nicolalde-Morejn et al. 1524, 1525 (XAL); Cihuat-ln,J. Borocio R. s.n.(ZEA); Cuautitln,Cochrane et al.10886(IBUG, WIS, ZEA),R. Cuevas et al 7025 (ZEA),

L. Guzmn & J. Santana M. 745, 947 (ZEA), F.Nicolalde-Morejn et al. 1528, 1529 (XAL), Prez dela Rosa 1039 (FCME, IBUG, MEXU), 1040 (FCME,IBUG, XAL), 1041 (CIB), 1518 (IBUG), Ramrez 425(IBUG),M. Rosales & L. Cruz 75(ZEA),J. Santana M.



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et al. 5296(ZEA); La Huerta,Cuevas et al. 4861(IBUG,ZEA); San Sebastin, F. Nicolalde-Morejn et al. 1422(XAL), 1423 (XAL), 1424 (XAL), 1425 (XAL), 1426(XAL), 1427 (XAL), Prez de la Rosa 1084 (IBUG,FCME, MEXU)1097, 1098(IBUG); Tuito,A. Flores etal. 614 (UAMIZ), F. Nicolalde-Morejn et al. 1429

(XAL), Prez de la Rosa 1438, 1439 (IBUG); Vallarta,Prez de la Rosa 1413, 1415(IBUG); Villa Puricacin,Prez de la Rosa 1885. (IBUG). NAYARIT: Gentry &Gilly 10496(MEXU),McVaugh 19211(FCME, MEXU),Vovides et al. 1487 (XAL), 1488 (XAL), 1489 (XAL),1490(XAL), 1491 (XAL), 1493(XAL); Compostela,F.

Nicolalde-Morejn et al. 1521 (XAL), 1522 (XAL),1523(XAL); Tepic,R. Dressler 1026(MO),H. S. Gentry& C. I. Gilly 10498 (FCME). Oaxaca: Miranda 4205(MEXU),F. Nicolalde-Morejn et al. 1465 (XAL), 1466(XAL), 1467 (XAL), 1468 (XAL), 1469 (XAL), 1470(XAL), 1471(XAL), 1472(XAL), 1473(XAL); Pochutla,Schutzman 543(XAL), 544(XAL), 545(XAL), 546(XAL),547(XAL), 548 (XAL), 550 (XAL), 551 (XAL), 552 (XAL),

553 (XAL), 554 (XAL), 555 (XAL), 556(XAL), 557(XAL),558 (XAL), 560 (XAL), 561 (XAL), 562 (XAL), 563 (XAL),565 (XAL),A. Nava-Zafra & J. Pascual 35 (SERO, FCME);Puerto Escondido, J. Rees 1603 (MO, XAL), Walters sn(FTG accession 714, XAL); San Gabriel Mixtepec, F.

Nicolalde-Morejn et al. 1474 (XAL); San Pedro Pochulta,J.Lomel et al. 2967(MEXU).

Zamia polymorpha D.W. Stev., A. Moretti& Vzq. Torres, Delpinoa n.s. 3738: 4.19951996 (issued 1998). Type: Belize.Cayo: 22 Jan 1989, D. W. Stevenson et al.

1119 (holotype: NY; isotypes: BRH, FTG,MO, NY, U). (Fig.10)

Stem hypogeous, branching dichotomouslywith age, up to 32 cm long, up to 14 cm indiam.Cataphylls chartaceous, persistent, basetriangular, apex aristate, 36.41.32.6 cm at

base, brown tomentose. Ptyxis inexed.Leaves 23(4), ascending, 301052945 cm, brown when emerging, green whenmature; petiole 10.295.3 cm long, greenish inyoung leaves, subterete, armed with pricklesup to 4 mm long; rachis subterete, up to 67 cmlong, with few prickles along the proximalthird. Leaets 312 pairs, sessile, coriaceous,lanceolate to oblanceolate, opposite to sub-opposite, apex acute, base attenuate, marginsserrulate along upper third, subrevolute; artic-ulations brown when young, green whenmature, 0.40.8 cm wide; the median leaets173523.5 cm.Pollen strobili usually 12,erect, conical, 6.57.3 cm long, 1.11.4 cm indiam, light to dark-brown tomentulose, apex

acute; peduncle light brown tomentose, 6.8 cmlong, 1.2 cm in diam; pollen sporangiophorescuneiform, distal face hexagonal-truncate,

0.4 cm long, fertile abaxial surface with 45bisporangiate synangia per lobe. Ovulatestrobili usually solitary, erect, cylindrical toovoid, 8.716.3 cm long, 58.3 cm in diam,dark-brown tomentulose, apex acute; peduncle

brown tomentose, 47.5 cm long, 1.1 cm indiam; megasporangiophores peltate, distal facehexagonal-truncate, 0.8 cm high, 1.82.1 cmwide. Seeds ovoid, sarcotesta pink whenimmature, red at maturity, 1.42.1 cm long,0.50.9 cm in diam, sclerotesta smooth.

Chromosome number.2n=17, 22, 23, 24,25, 26, 27, 28 (Stevenson et al.,19951996b,

b; Vovides & Olivares,1996).Distribution and habitat.This species is

known from Belize, Guatemala and Mexico

from 0

200 m elevation. In Mexico, its rangeincludes the states of Quintana Roo, Yucatn,Campeche, Tabasco and Chiapas (Fig.6).

Etymology.The specic epithet alludes tothe extreme variation in leaf and leaet mor-

phology presented by this species (Stevensonet al., 19951996b).

Distinguishing features.Zamia polymor-pha shares many morphological attributeswith Z. loddigesii of the Gulf of Mexicodrainage. However, there are clear differences

in the pollen and ovulate reproductive struc-tures. In Z. loddigesii, the pollen strobili arebeige in color with an acute apex and theovulate strobili are beige and cylindricalwhereas in Z. polymorpha the pollen strobiliare maroon with an acuminate apex and theovulate strobili are dark-maroon and ovoid.

Additional specimens examined.MEXICO.CAMPECHE: Hernndez et al. ES-184 (MEXU), Schutz-man 502(XAL), 503(XAL), 504(XAL), 505(XAL), 506(XAL), 507 (XAL), 508 (XAL), 509 (XAL); BenitoJurez, Vovides et al. 1312 (XAL), 1313 (XAL), 1314(XAL), 1315(XAL), 1316(XAL), 1317(XAL); Calakmul,

Madrid et al. 736 (MEXU), Martnez 30420-A (MEXU);Ciudad del Carmen,Flores et al. 9586(XAL); Champoton,Chan 3719(CICY, XAL),Vovides 853(XAL), 854(XAL),855 (XAL), 1326 (XAL), 1328 (XAL), 1329 (XAL),1330(XAL), 1331(XAL), 1332(XAL), 1333(XAL), 1334(XAL), 1335 (XAL), 1336 (XAL), 1337 (XAL), 1338(XAL), 1339(XAL), 1527(XAL); Hopelchen,Ortega &Ucn 1562 (UADY, XAL), Ucn et al. 7293 (UADY,XAL), 7307 (UADY, XAL), 7398 (UADY, XAL).CHIAPAS:Palenque,Aguilar & Aguilar 1355 (MEXU),F.

Nicolalde-Morejn & N. Martnez 1419 (XAL),Schutzman508 (XAL), Walters s.n. (FTG accession 13-2, XAL);

Ocosingo, Walters s.n. (FTG accession 17-2, XAL).QUINTANAROO:Cabrera et al. 2574(MEXU),Davidse etal. 20075(MEXU, MO),Tllez 1415(MEXU),Trejo 225(CICY, MEXU); Adolfo Huerta, lvarez et al. 9495



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(MEXU); Chetumal,Vovides 852(XAL),Flores & Burgos9635(XAL), 9643(XAL), Othon P. Blanco,Vovides et al.1318 (XAL), 1319 (XAL), 1320 (XAL), 1321 (XAL), 1322(XAL), 1323(XAL), 1324(XAL), 1325(XAL). TABASCO:Balancan, Matuda 3117 (MEXU), Mndez 214 (XAL),

Novelo 169 (MEXU, XAL), Puig 788 (MEXU). Macu-spana, Vovides et al. 1344, 1345 (XAL). Yucatn, G. F.Gaumer 2430(MO), Lundell & Gentle 827(MEXU),May743 (CICY, MEXU); Tekon, Enrquez 94 (MEXU);

Tzucacab,Vovides et al. 1303 (XAL), 1306(XAL), 1307(XAL), 1308 (XAL), Flores & Burgos 9642 (XAL);Valladolid, Vovides 856, 857 (XAL), Vovides et al. 867(XAL), 868(XAL), 869(XAL) 871(XAL), 872(XAL),873 (XAL), 874 (XAL), 875 (XAL), 881 (XAL), 877

(XAL), 870 (MEXU, XAL), 876 (MEXU, XAL), 877(MEXU, XAL), 878(MEXU, XAL), 880 (MEXU, XAL);Yaxcaba, Vovides et al. 1309(XAL), 1310(XAL), Vovides1311(XAL).

FIG. 10. Zamia polymorpha. A. Habit (Stevenson et al. 1118, NY).B. Leaets (from the holotype). C. Ovulatestrobilus.D. Pollen strobilus.E. Cataphyll.



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GUATEMALA. PETEN: W. E. Harmon & J. A.Fuentes 5735(MO).

BELIZE. BELIZEDISTRICT:Estrada 234(CICY),D.L. Spellman 1548 (MO), C. Whitefoord 2603 (MO).CAYODISTRICT:D. W. Stevenson et al. 1121 (FTG, MO,

NY, U), 1122 (FTG, MO, NY, U), M. J. Balick et al.1803 (MO, NY), M. J. Balick et al. 2058 (NY), T. B.Croat 23732 (MO), D. R. Hotel & L. Thomas 1130(NY),J. S. Huston s.n.(MO),R. W. Long 3238(MO),J.

A. Ratter 5195 (MO),D. L. Spellman 1974(MO),D. L.Spellman & W. W. Newey 1962 (K), J. R. Wiley 333(MO). Orange Walk, G. Davidse & A. E. Brant 32768(MO). Stann Creek, R. L. Walter 1099 (MO). Sin datos(K).

Zamia prasina W. Bull, Retail List: 20.1881. Type: cultivated from Br. Honduras,W. Bull s.n. (holotype: K).

Stem hypogeous to epigeous, rarelybranching.Cataphyllschartaceous, persistent,base triangular, apex aristate, 34.51.22.7 cm at base, brown tomentose. Ptyxisinexed. Leaves 24(6), 571002535 cm,ascending to spreading, brown when emerg-ing; petiole 1230 cm long, brown-greenishyoung leaves, subterete, sparsely to denselyarmed with prickles up to 4 mm long; rachissubterete, up to 70 cm long, with few prickles

along the proximal third. Leaets 12

18pairs, sessile, coriaceous, oblong to oblanceo-late, opposite to subopposite, apex acute toacuminate, base cuneate, margins serrate todenticulate in the upper two third, subrevo-lute; articulations brown-yellowish in youngleaets, 0.30.6 cm wide; the median leaets152046 cm. Pollen strobili usually 12,erect, cylindrical to ovoid, up to 610 cmlong and 24 cm in diam, light brown, apexacute; peduncle densely light brown tomen-tose, 24 cm long, 1.5 cm in diam. Ovulate

strobiliusually solitary, erect, cylindrical, 1015 cm long, 57 cm in diam, green, glabrouswhen mature, apex acute; peduncle brown-greenish tomentose, 3.5 cm long, 1.5 cm indiam; megasporangiophores distal face hex-agonal-truncate, 11.4 cm high, 1.11.8 cmwide. Seeds ovoid, sarcotesta light red whenimmature, red at maturity, 1.52 cm long,0.50.8 cm in diam, sclerotesta smooth.

Chromosome number.Not known.Distribution and habitat.Endemic to

Belize (Fig. 6) on rocky outcrops between100200 m elevation in evergreen tropicalrainforest.

Etymology.The specic epithet alludes tothe bright grass-green leaets.

Distinguishing features.Plants with sub-terranean or epigeal stems, distinctly serrulateleaet margins and bright-green leaets;

ovulate strobili ovate, green and glabrouswhen mature.

Additional specimens examined.BELIZE.TOLEDO: G. Davidse & A. E. Brant 32179 (MO),32232(MO).

Zamia purpureaVovides, J.D. Rees & Vzq.Torres, Flora de Veracruz 26: 28. 1983.Type: Mexico. Veracruz: 30 Jan 1982,Vovides 734(holotype: XAL). (Fig.11)

Stemhypogeous, dichotomously branchingwith age, up to 30 cm long, 46 cm in diam.Cataphylls membranaceous, deciduous, basetriangular, apex acuminate, 5.1.8 cm at

base, reddish-brown tomentose. Ptyxis in-exed. Leaves 16, ascending to spreading,34903238 cm, reddish-brown whenemerging, turning green to dark-green atmaturity; petiole 1629 cm long, blackish inyoung leaves, subterete, armed with simple

prickles up to 4 mm long; rachis subterete,1745 cm long, with few prickles along theproximal third.Leaets 34(6) pairs, sessile,coriaceous, elliptic to lanceolate, opposite tosubopposite, veins highly prominent on ad-axial surface, though relatively inconspicuousveins have been reported on some individu-als, apex acute, base attenuate, marginsdentate along upper third, subrevolute; artic-ulations brown in young leaets, 0.40.8 cmwide; the median leaets 62728.1 cm.

Pollen strobiliusually 12, erect, conical, 24.3 cm long, 0.51.1 cm in diam, light-browntomentulose, apex acute; peduncle light-

brown tomentose, 23.5 cm long, 0.70.9 cm in diam; pollen sporangiophorescuneiform peltate, distal hexagonal face andtruncate to scutiform, 0.25 cm long, fertileabaxial surface with 2 bisporangiate synangia

per lobe. Ovulate strobili usually solitary,erect, conical, 69 cm long, 34.5 cm indiam, purplish-brown tomentulose when im-mature turning dark-purple glabrescent when

mature, apex acute; peduncle densely dark-brown tomentose, 2.9 cm long, 1.6 cm indiam, distal face hexagonal-scutiform, 0.9



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FIG. 11. Zamia purpurea.A. Habit. B. Pollen strobilus.C. Microsporophyll, abaxial view.D. Ovulate strobilus.E.Ovulate sporangiophores.F. Seed. (FromRees1 654, XAL.)



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1.2 cm high, 1.62.1 cm wide. Seeds ovoid,sarcotesta pink when immature, red at matu-rity, 0.81.1 cm long, 0.60.8 cm in diam,sclerotesta smooth.

Chromosome number.2n= 16 (Vovides,

1983)Diversity and genetic structure.The

average of alleles per locus is A=2.10, thepercentage of polymorphic loci is P= 100,the expected heterozygosity is HE=0.481 andthe genetic differentiation between the two


Distribution and habitat.Zamia pur-purea is endemic to Mexico from the RoUxpanapa drainage system within the states

of Veracruz and Oaxaca (Fig.5) between 50

200 m in evergreen tropical forest (sensuRzedowski,1978) classication.

Etymology.The specic epithet alludes tothe dark purple color of the ovulate strobili.

Distinguishing features.Characterized bythe prominent leaet veins giving the coria-ceous leaets a channeled appearance incontrast to the rest of its congeners inMesoamerica, and the dark purple color ofthe ripe ovulate strobili.

Additional specimens examined.MEXICO.OAXACA: Santa Mara Chimalapa, S. H. Salas M. 982(SERO), Snchez et al. 40 (B, MEXU); Santa MaraLachixio,Cern et al. 266(XAL); San Juan Guichicovi,F.

Nicolalde-Morejn & J. Torres 1503(XAL), 1404 (XAL),M. Vzquez-Torres et al. 1470 (MO, XAL), Walters s.n.(FTG accession 101, XAL). VERACRUZ: M. Vzquez-Torres 4038 (CIB); Hidalgotitln, Calzada 8374 (XAL),

M. Vzquez-Torres et al. 224 (MO); Jess Carranza, A.Gonzlez-Christen s.n. (CIB), F. Nicolalde-Morejn & J.Torres 1502(XAL), M. Vzquez-Torres et al. 2373 (CIB),

M. Vzquez-Torres et al. V-2532(CHAPA, CIB, XAL).

Zamia sandovalii C. Nelson, Ceiba 46(12):55.. 2005. Type: Honduras. Atlntida: Jan2006, G. Sandoval et al. 1154 (holotype:TEFH)

Stem hypogeous, non-branching, 15.7 cmlong, 10.1 cm in diam. Cataphylls persistent,

base triangular. Leaves 13, ascending todescending up to 2100.55 cm; petiole 68 cmlong, subterete, armed with small prickles;rachis subterete, up to 140 cm long, unarmed.

Leaets68 pairs sessile, sub-coriaceous, lance-olate, opposite to subopposite, falcate, apexacuminate, base attenuate, margins serrulate

along distal third, subrevolute; the median leaf-lets 163123 cm. Pollen strobili usually 2,erect, cylindrical, 11.2 cm long, 2.6 cm in diam,

brown-reddish tomentulose, apex apiculate; pe-duncle light-brown tomentose, 16.7 cm long,

1.2 cm in diam; pollen sporangiophores cunei-form, distal face hexagonal truncate, 0.8 cmlong, fertile abaxial surface with 23 bisporan-giate synangia per lobe.Ovulate strobiliusuallysolitary, erect, ellipsoid to cylindrical, 12 cmlong, 4.5 cm in diam, brown-yellowish, tomen-tulose, apex long-acuminate; peduncle densely

brown tomentose, 1.5 cm long; megasporangio-phores peltate, distal face hexagonal-truncate,22.5 cm high, 1.82 cm wide. Seeds ovoid,sarcotesta white when immature, 1.21.8 cm

long, 0.5

0.7 cm in diam, sclerotesta smooth.Chromosome number.UnknownDistribution and habitat.Endemic to

Honduras (Fig. 6), between 200350 m inevergreen tropical forest.

Etymology.The specic epithet honorsGermn Sandoval, biologist of the Universidad

Nacional Autonoma de Honduras (Nelson,2005).

Distinguishing features.This species ischaracterized by glabrous, sub-coriaceous,

lanceolate, falcate lea

ets; cylindric, ovulatestrobili with a strongly acuminate apex; andseeds with a white sclerotesta.

Additional specimens examined.HONDURAS.ATLNTIDA: J. Haynes et al. 37 (TEFH), G. Sandovalet al. 1155 (TEFH),1156(TEFH).

Zamia soconuscensisSchutzman, Vovides &Dehgan, Bot. Gaz. 149(3): 347.1988. Type:Mexico. Chiapas: Feb 1939, Matuda 2659(holotype: F; isotypes: CR, MEXU, MICH).

Stem epigeal, erect to decumbent in adultplants, branching dichotomously with age,3065 cm long, 1031.5 cm in diam. Cata-

phyllschartaceous, persistent, base triangular,apex aristate, 7.12.6 cm at base, reddish-

brown tomentose. Ptyxis inexed to erect.Leaves315 or more per apex, 1201904562 cm, ascending, distal portion descendingto spreading, brown when emerging turninggreen at maturity; petiole 3872 cm long,

green-yellowish in young leaves, terete,armed with prickles up to 5 mm long; rachissubterete, up to 84 cm long, with few prickles



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along the proximal third. Leaets 4152pairs, sessile, coriaceous, linear-lanceolate,alternate to subopposite, subfalcate, apexacute, base attenuate; margins entire, subre-volute; articulations brown in young leaets,

0.40.8 cm wide; the median leaets 12350.61.5 cm. Pollen strobili usually 13 perapex, erect, cylindrical to conical, 915 cmlong, 1.22.4 cm in diam, dark-brown tomen-tulose, apex apiculate; peduncle light-browntomentose, up 7.2 cm long, 1.2 cm in diam;

pollen sporangiophores cuneiform, distal facehexagonal-truncate, 0.45 cm long, fertileabaxial surface with 5 bisporangiate synangia

per lobe. Ovulate strobili usually solitary,erect, cylindrical, 1215 cm long, 6.17.3 cm

in diam, dark-brown to reddish, tomentulose,apex aristate; peduncle brown puberulent,2.1 cm long, 1.2 cm in diam; megasporan-giophores peltate, distal face hexagonal-truncate, 0.70.9 cm high, 1.61.8 cm wide.Seeds ovoid to angular, sarcotesta white whenimmature turning salmon-pink when mature,sclerotesta light-beige, smooth with 68 lightfurrows running longitudinally and some-times dichotomizing, 2.12.6 cm long, 1.41.9 cm diam.

Chromosome number.

2n=16 (Schutzmanet al.,1988).Distribution and habitat.Endemic to

Chiapas, Mexico, between 9001400 m inthe Soconusco mountain range of southernChiapas. It inhabits the understory herbaceouslayer of the transition zone between evergreentropical forest and cloud forest (Fig.5).

Etymology.The specic epithet alludes tothe Sierra del Soconusco mountain range alsoknown as the Sierra Madre de Chiapas, beingthe region where this species is native(Schutzman et al., 1988).

Distinguishing features.Zamia soconus-censis is the only species of the genus inMexico that approaches an arborescent habitwith leaves that can reach almost two meterslong that gracefully arch toward the terminal

portion. The linear-lanceolate leaets havetotally entire margins; the only other Mexicancongener with entire margins is Z. inermis.Ovulate strobili are short-pedunculate and sol-itary with a dark-brown velvety tomentulum.

Additional specimens examined.MEXICO.CHIAPAS: Matuda 2087(MEXU), 2535 (MEXU), 2590

(MEXU), 2656 (MEXU); Acacoyagua, Garca 149(CHIP, MEXU), M. A. Prez-Farrera 141 (CIB, HEM,MEXU).

Zamia spartea A. DC., Prodr. 16 (2): 539.

1868. Zamia loddigesii var. spartea (A.DC.) Schuster, Panzenr. 99: 148. 1932.Type: Mexico. Oaxaca: prope Acayucam,Verapa, Chimalapi, 1832, Alaman s.n.(holotype: G-DC). (Fig.12).

Stemhypogeous, branching dichotomouslywith age, 540 cm long, 58 cm in diam.Cataphyllschartaceous, persistent, base trian-gular, apex aristate, 6 1.4 cm at base,yellowish tomentose. Ptyxis inexed. Leaves

2

5(8) per crown, 35

6038

52 cm, ascend-ing to gracile, reddish-brown when emerging,turning green at maturity; petiole 1221 cmlong, green-yellowish in young leaves, sub-terete, heavily armed with straight to some-times bifurcate prickles up to 4 mm l

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Revision of zamia in mega mexico